Review of Acantholaimus Allgén, 1933 (Nematoda: Chromadoridae), a genus of marine free-living nematodes, with a tabular key to species

in Nematology
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Acantholaimus (Nematoda: Chromadoridae) is considered to be one of the most abundant and species-rich genera of deep-sea free-living nematodes and has a world-wide distribution. To date, 50 Acantholaimus species are known globally, although in all the oceans many more undescribed species have been recorded. Local species diversity of the genus is often very high and may amount to several dozen species in one deep-sea locality. The present review summarises data on morphology (with illustrations), taxonomy, ecology, and biology of this genus. An emended genus diagnosis, a list of species with their known locations, and a tabular key to species are provided. Acantholaimus ewensis Platt & Zhang, 1982 n. syn. is proposed as a junior synonym of A. polydentatus Gerlach, 1951.

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References
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Figures
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    General appearance of Acantholaimus males, line drawings from light microscope. A: A. cornutus; B: A. occultus; C: A. veitkoehlerae; D: A. angustus. (Scale bars = 100 μm.) From Miljutina & Miljutin (2012).

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    Light micrographs of semi-thin cross-sections of Acantholaimus specimens. A: Cross-section at level of cheilostom; B: Cross-section at level of pharynx; C: Cross-section at level of intestine; D: Cross-section of spermatozoon at level of male reproductive system; E: Cross-section of proximal end of male spicules; F: Cross-section of male spicule at distal end (cloacal region). Abbreviations: d.ch. = dorsal hypodermal chord; gu. = gubernaculum; i. = intestine; l.ch. = lateral chord; l.f. = lateral field of cuticle; ph. = pharynx; ru. = ruga; sd.ch. = subdorsal hypodermal chord; sv.ch. = subventral hypodermal chord; sp. = spicule; sz.cy. = spermatozoon cytoplasm; sz.nl. = spermatozoon nucleolus; sz.nu. = spermatozoon nucleus; v.ch. = ventral hypodermal chord. (Scale bars = 10 μm.)

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    Different arrangements of dots in the cuticular lateral field seen in Acantholaimus species, light microscope photographs. A: Longitudinal rows; B: No distinguishable differentiation; C, D: Dots larger and arranged irregularly compared to remaining cuticle. (Scale bars = 10 μm.)

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    Head and cervical sensilla in different Acantholaimus specimens, SEM photographs. A, B, D: Specimens with pairs of closely grouped anterior cervical setae and relative small amphids; C: Specimen with numerous closely grouped anterior cervical setae; E: Specimen with paired anterior cervical setae and large amphid. (Scale bars = 10 μm.)

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    Acantholaimus sp., a head, front view. SEM photograph. (Scale bar = 10 μm.)

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    Heads of different Acantholaimus species, line drawings from light microscope. A: A. formosus; B: A. robustus; C: A. veitkoehlerae; D: A. angustus; E: A. cornutus; F: A. tchesunovi; G: Everted head of A. veitkoehlerae, front view; H: A. sieglerae; I: A. quintus; J: A. arthrochaeta. Abbreviations: a. = amphid; l = labium; m.o. = mouth opening; r. = ruga; t. = onchium. Illustrations taken from Miljutina & Miljutin (2012) (B-H, J) and Miljutina et al. (2013) (A, I). (Scale bars = 20 μm.)

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    Somatic setae and pores in different Acantholaimus specimens, SEM photographs. A, B: Mid-body region; C, D: Cervical region. Large pores that appear to represent pits of lost somatic setae are marked with arrows. (Scale bars = 10 μm.)

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    Acantholaimus robustus, head. Reconstruction based on micrographs made using a confocal microscope. Abbreviations: a. = amphid; l. = lip; r. = cuticular ruga in cheilostom; t. = onchium. (Scale bar = 20 μm.) From Miljutina & Miljutin (2012).

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    Anterior end of Acantholaimus species, line drawings from light microscope. A: A. veitkoehlerae; B: A. barbatus; C: A. cornutus. From Miljutina & Miljutin (2012). (Scale bars = 50 μm.)

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    Acantholaimus sp., male spicules and gubernacula, SEM photographs. A: Distal end of spicules; B, C: Distal end of gubernacula. (Scale bars: A = 10 μm; B, C = 1 μm.)

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    Construction of male spicules and gubernacula, female reproductive system and tail in Acantholaimus species, line drawings from light microscope. A-I: Spicules and gubernacula of A. arthrochaeta (A), A. maks (B), A. cornutus (C), A. seiglerae (D), A. tchesunovi (E), A. barbatus (F), A. robustus (G), A. occultus (H), A. veitkoehlerae (I); J: Female reproductive system of A. angustus; K: Posterior end of A. veitkoehlerae. From Miljutina & Miljutin (2012). (Scale bars: A-I = 10 μm; J, K = 50 μm.)

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    Contents of intestine in Acantholaimus specimens, light micrographs. A, C: Mixture of fine organic and inorganic particles; B: Large unicellular organisms; D: Remnants of unicellular organisms with thick body walls (?). (Scale bars = 20 μm.)

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    Acantholaimus specimen with a scarred stump just posterior to anus, light micrographs. A: View of scarred stump; B: Food particles in intestine indicating active feeding of the injured specimen. (Scale bars = 20 μm.)

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